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Calopterygidae
	Immature adult male Calopteryx virgo
	Adult male Calopteryx virgo
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Odonata
Suborder:	Zygoptera
Superfamily:	Calopterygoidea
Family:	Calopterygidae; Sélys, 1850
Subfamilies
	Caliphaeinae; Calopteryginae; Hetaerininae; See text for genera

Calopterygidae is a family of damselflies, in the suborder Zygoptera.^[2] They are commonly known as the broad-winged damselflies,^[3] demoiselles, or jewelwings.^[4] These rather large damselflies have wingspans of 50–80 mm (compared to about 44 mm in the common bluetail damselfly, Ischnura elegans), are often metallic-coloured, and can be differentiated from other damselflies by the broader connection between the wings and the body, as opposed to the abrupt narrowing seen in other damselfly families.^[5] The family contains some 150 species.

The Calopterygidae are found on every continent except Antarctica. They live along rivers and streams.^[6]

Etymology

The name is derived from Greek kalos meaning beautiful and ptery meaning winged.

Characteristics

The adults have metallic bodies; their wings are broader, with wider bases than other damselflies, and at rest hold their wings parallel to the body, slightly elevated. Some species have conspicuously colored wings; in males, the wings are usually blue, without pterostigmata, in females green or brown.^[7]^[8] Species are often quite variable in color and patterning, and they are sexually dimorphic. Color intensity may fade with age.^[6] The wings are heavily veined, having often 18 or more antenodal veins. The first segment of their antennae is longer than the combined length of the other segments. They have a jerky, skipping form of flight similar to the flight pattern of a butterfly (fluttering, rather than hovering stably like many other damselfly and dragonfly families). At least one species of Calopterygiadae has shown morphological plasticity in wing length due to the closeness of a forest to the river or stream where they live with a further forest correlating to greater wing length.^[9] They perch horizontally on twigs near the water's edge.^[7]^[8]

Calopterygidae nymphs have lateral gills are longer than the median gills.^[10] The nymphs have a flattened, pentagonal-shaped head, a long first antennal segment and long legs. They are found among submerged aquatic plants, woody debris and the exposed roots of streamside plants. There is a single generation per year.^[8] The time spent in the larval stage is influenced by both biotic factors, such as fat reserves, and abiotic factors, such as temperature, so they have the highest chances of surviving and reproducing.^[11]

Behavior

Hetaerina americana mating: sperm removal

The mating system of most species in this family is resource defense polygyny, where males are often territorial, guarding riverine habitat that is sought after by females for egg deposition. Some males are not territorial. Within a species there may be a territorial and nonterritorial morph, which may be different in coloration.^[6]

Some species display courtship behavior, especially displays of wing movement by the male.^[6] At least one genus (Hetaerina) displays lekking behavior.^[12]

During mating, the male first removes other males' sperm from the female's reproductive tract, then places his own sperm there. The intromittent organ of the male has spines that physically remove rival sperm and also stimulate the female's muscles to contract and expel the sperm. In many species, the male accompanies the female when she searches for a site to lay eggs; in some cases, he even remains attached to her.^[6] The guarding of females post-copulation is done so another male does not mate with the female before laying her eggs even though the male may be able to reproduce with other females and in the case of Hetaerina species, the male may lose his territory during the time spent guarding.^[13]

Like all Odonata species, the species in this family are carnivorous in both their larval and adult stages. Larvae tend to feed on smaller invertebrates, such as mayflies.^[10]

Classification

Subfamiles and tribes according to Dijkstra et al. (2014)^[14]

Subfamily Calopteryginae Selys, 1859 – the demoiselles:

Tribe Caliphaeini Fraser, 1929
- Caliphaea Hagen, 1859
Tribe Calopterygini Selys, 1850
- Atrocalopteryx Dumont, Vanfleteren, De Jonckheere, & Weekers, 2005
- Calopteryx Leach, 1815
- Matrona Selys, 1853
- Matronoides Foerster, 1897
- Neurobasis Selys, 1853
Tribe Iridictyonini Dumont et al., 2005
- Iridictyon Needham & Fisher, 1940
Tribe Mnaisini Ishida, 1996
- Archineura Kirby, 1894
- Echo Selys, 1853
- Mnais Selys, 1853
- Psolodesmus McLachlan, 1870
Tribe Noguchiphaeini Dumont et al., 2005
- Noguchiphaea Asahina, 1976
Tribe Saphoini Dumont et al., 2005
- Phaon Selys, 1853
- Sapho Selys, 1853
- Umma Kirby, 1890
Tribe Vestalini Needham, 1903
- Vestalaria May, 1935
- Vestalis Selys, 1853

Subfamily Hetaerininae Selys, 1853 – the rubyspots and others:

Bryoplathanon Garrison, 2006
Hetaerina Cowley, 1934
Mnesarete Hagen in Selys, 1853
Ormenophlebia Garrison, 2006

References

^ Selys-Longchamps, E. (1850). "Revue des odonates ou libellules d'Europe". Mémoires de la Société Royale des Sciences de Liège (in French). 6: 1-408 [133] – via Biodiversity Heritage Library.
^ "Family CALOPTERYGIDAE". Australian Faunal Directory. Australian Biological Resources Study. 2014. Retrieved 30 March 2017.
^ Calopterygidae. Digital Key to Aquatic Insects of North Dakota. Valley City State University.
^ Calopterygidae. Archived 2016-10-05 at the Wayback Machine Dragonflies and Damselflies of Ecuador. Electronic Field Guide Project, University of Massachusetts, Boston.
^ “Broad-Winged Damselflies.” Calopterygidae Family - Broad-Winged Damselflies, https://unsm-ento.unl.edu/Odonata/calo.html.
^ ^a ^b ^c ^d ^e Córdoba-Aguilar, A. & Cordero-Rivera, A. (2005). Evolution and ecology of Calopterygidae (Zygoptera: Odonata): status of knowledge and research perspectives. Neotrop. Entomol 34(6), 861-879.
^ ^a ^b Dijkstra, K. B. Field Guide to the Dragonflies of Britain and Europe. British Wildlife Publishing, 2006. ISBN 0-9531399-4-8. Pages 23, 65.
^ ^a ^b ^c Capinera, J. L. (2008). Encyclopedia of Entomology. Springer Science & Business Media. pp. 1243–1244. ISBN 978-1-4020-6242-1.
^ Taylor, Philip D., and Gray Merriam. “Wing Morphology of a Forest Damselfly Is Related to Landscape Structure.” Oikos, vol. 73, no. 1, May 1995, p. 43., https://doi.org/10.2307/3545723.
^ ^a ^b Calopterygidae. UNH Center for Freshwater Biology.
^ Córdoba-Aguilar, Alejandro, and Adolfo Cordero-Rivera. “Evolution and Ecology of Calopterygidae (Zygoptera: Odonata): Status of Knowledge and Research Perspectives.” Neotropical Entomology, vol. 34, no. 6, Dec. 2005, pp. 861–879., https://doi.org/10.1590/s1519-566x2005000600001.
^ Córdoba-Aguilar, A., et al. (2009). The lek mating system of Hetaerina damselflies (Insecta: Calopterygidae). Behaviour, 146, 189-207.
^ Alcock, John. “Post-Copulatory Mate Guarding by Males of the Demselfly Hetaerina Vulnerata Selys (Odonata: Calopterygidae).” Animal Behaviour, vol. 30, no. 1, Feb. 1982, pp. 99–107., https://doi.org/10.1016/s0003-3472(82)80242-x.
^ Dijkstra, Klaas-Douwe B.; Kalkman, Vincent J.; Dow, Rory A.; Stokvis, Frank R.; et al. (2014). "Redefining the damselfly families: a comprehensive molecular phylogeny of Zygoptera (Odonata)". Systematic Entomology. 39 (1): 68–96. Bibcode:2014SysEn..39...68D. doi:10.1111/syen.12035.